Documente Academic
Documente Profesional
Documente Cultură
PART III
VEGETA TI
'10
Api seedling plant undergo shifts in their "ontogenetic age" as cells in their apical meristem divide and lay down new tis sue. Axillary buds typically are dormant due to apical dominance, but retain lhe ontogenetic age at origino Once lateral buds start to grow they then undergo change in maturation in lhe same manner as the apical bud, but in their own pattem.
FIGURE 16-15
Apical Meri
eovers a mutation (79). This situation also oeeurs in the budding offruit and nut tree cultiv Although the probability for any single mutation to oceur may be low, the probability of ing it is high because of the single bud propagation and the volume of plants involved. T; explains why off-type plants are usually detected in a commercial orchard rather than in -source orehard where the propagation material was obtained. li the eharacter affects flo ing or fruiting, the mutant may go undetected for a number of years after the plant is pl into its permanent location.
Homoblastic phase changes that involve little obvious change in phenotype with development and age.
Heteroblastic phase changes that show distinct change in phenotype during development.
CHAPTER 16
609
GETTING
MORE
IN DEPTH
ON THE
SUBJECT
Phase changes can be considered to be biologically equivalent in animals, humans, and plants in that embryonic, juvenile, adolescent, and adult phases are present in each. The difference is that animals and plants differ in the way they grow and age. Cells in animal (and human) development are more or less determinate (i.e., cells in all parts of the body mature and age more or less together). Plants, on the other hand, grow from consecutive cell divisions in the meristem of apical and lateral shoots laid down in sequence over time. Juvenile to mature change occurs in the apical
meristem as the shoot grows, but cells remaining behind retain their initial ontogenetic age. In perennials, ontogenetic aging continues in altemating cyc1es of grawth and dormancy. ln annuals and biennials it terminates in reproductive structures and the plant dies. The phase of maturation in different locations of the plant is deterrnined by the pattem of bud development in the plant. Juvenile and mature characteristics, such as flowering, leaf shape, or thorniness, can be used as markers of maturation, their location depending on the chronological age at which ontogenetic maturity is attained in particular parts of the plant.
GETTING
MORE
IN DEPTH
ON THE
SUBJECT
The comparison of the two types of "aging" produces a paradox in horticultural terminology. That part of a seedling plant nearest to its base is the "oldest" in terms of chronology but actually "youngest" (i.e., more juvenile) in terms of maturity (onto-
Paradox in Terminology
genetic) age. Likewise, the outer peripheries of the stems and branches are the "oldest" in maturity but "youngest" in chronology (15, 35, 74). The age of a vegetatively prapagated plant depends on the ontogenetic age at which the plant was prapagated.
have been identified that are differentially expressed in the juvenile or mature. These can be used as molecular or biochemical markers of phase development (57). Chronological aging continues through the life of an individual plant whether seedling or vegetatively prapagated. This characteristic may best be designated by the number of years that the plant has grown either fram seed or vegetative prapagule. When young and growing under favorable conditions, the plant should be vigorous, healthy, and may become very floriferous, Eventually, as the plant gets older, vigor declines and, although flowering and fruiting may remain abundant, new growth will tend to be reduced. The plant eventually becomes enescent and may eventually die. Plants appraaching the senescent stage can be returned to vigorous and productive state by reprapagation or by applying various horticultural practices, such as pruning, nitragen fertilization, irrigation, controlling pests and diseases, and, in general, applying good management practices.
Juvenile phenotype characteristics associated with the juvenile phase in specified species, usual! y upright growth, vigor, sometimes thorniness, and lack of flowering.
610
PART III
Adult phenotype characteristics associated with the adult phase, typically bushy growth, reduced vigor, no thorns, and profuse f1owering.
of the plant tend to produce progeny that are biologically mature; the entire range may be produced if plants from intermediate locations are included. The recognition of this variabili . phenomenon by foresters led to the coining of the terms ortet (the So plant) and ramet ( plants) (138). These variations are not due to genetic change but to epigenetic variation in maruration that persists during vegetative propagation.
"Bulking up" in forestry propagation this term refers to using vegetative propagation to multiply the supply of se!ected genotypes.
CHAPTER 16
PRINCIPLES
GETTING
MORE
IN DEPTH
ON THE
SUBJECT
(a) Seed is taken from plants of seedling families that have been previously selected for superior growth rates, form, disease resistance, and so on. (b) These seedling blocks (referred to as ortets) are kept in a juvenile/transition phase by hedging and shearing. (c) Cut(ramets) are obtained and can be rooted because of
their juvenile condition. (d) Rooted plants are transplanted to timber production sites for evaluation over time. (e) Superior ortet farnilies are consequently identified. (f) Cuttings, taken from the seedling plants of these selected elite families which have been maintained by hedging and shearing are multiplied into commercial volumes ("bulking up"). (g) Clonally generated ramets are established in the forest planting in a mosaic of different clones to avoid monoculture production and to reinforce genetic diversity.
PART m
Graph showing the effect of age on donor tree relating to key tree characteristics that are important in cJonal forestry. Left: Phenotypic trueness-totype is illustrated by size of stem diameter and shoot length of grafts after multiple years. Right: rooting percentages of cuttings from donor trees of different ages. (From Greenwood et aI., in Clonal Forestry 11, 1993 by permission.)
E ...
6 5 100
C1) C1)
.S! ~
c:
:g
c
(.J
.s
c: :;::
CI
....
40
Eastern Larch
... ...---t)6
~ o
..... E (.J
20
30
40
50
00
20
40
60
Age (years)
"Clonal forestry" depends on the ability to mass-propagate populations of juvenile prapagules that not only exhibit easy rooting (Figure 16-18, right), but also praduce the desirable juvenile "seedling'' phenotype (Figure 16-18, left) (2). The desirable "true-to-type" form of forest trees grown for lumber and wood is straight-tapered trunk, reduced branching, lateness to flower, and other juvenile characteristics. Vegetatively prapagated plants of radiata pine (Pinus radiata), loblolly pine (Pinus), giant sequoia (Sequoiadendron), and eastern larch (Larix) taken fram more mature locations in the tree showed reduced height and diameter, and more branching (53). Success in "clonal forestry" depends on the ability to contrai maturation and to select the proper "true-to-type" stage (2, 116). See the text box on page 611 (24, 116, 129).
CHAPTER 16
613
Epicormic a shoot emerging from a latent bud on the base of a tree; synonymous with watersprout.
o
Cone of juvenility a cone-shaped area comprising the trunk and lower branches of a seedling tree, which tends to remain juvenile.
Ontogenetic juvenile-mature gradients occur in seedling trees from the base of the tree to the topo Left: Seedling conifer. Juvenile root-shoot junction occurs at A; flowering occurs first at the mature apex B. Other letters refer to shoots at different locations. Gradient in juvenility = A> B > E > D > C > F due to differences in ontogenetic age. Right: Seedling deciduous tree. Juvenile root-shoot junction occurs at A; flowering occurs first in the mature part of tree at the apical part (G). Juvenile structures arising from the "cone of juvenility" near the base of the tree include adventitious root "sucker" (B), watersprout (epicormic) (C), and sphaeroblast (D). A stump sprout from pruning (E) and a hedged tree (F) are also shown. (Redrawn with permission from Bonga [16].)
FIGURE 16-18
Stump sprouts vigarous shoots that are produced from the stump when a tree is pruned back severely.
Sphaeroblasts masses of adventitious buds produced on the lower trunk of some tree species.
Hedge row a row of trees or bushes pruned back to a hedge to stimulate shoots for propagation.
Serial propagation propagation of seedling ar clonal material for several consecutive generations in containers, each propagated after pruning.
614
PART III
VEGETATNE
PROPAGA TIO
Mature shoot
Juvenile shoots
Rejuvenation produced by consecutive (serial) grafting from a ten-year-old mature Eucalyptus tree. Left: Plant produced by cleft graft of scions from the ten-year-old tree onto a seedling rootstock showed mature leaves. Middle: Plant produced from scions after three serial grafis. One of the shoots shows mature characteristics, the other juvenile. Right: Plant produced after grafiing with scions afier six serial grafts. AlI of the shoots produced juvenile (large) leaves. Cuttings from this plant rooted successfully, whereas rooting from the ten-year-old original mature tree was not successful. (From Sinisca1co and Pavolettoni (126) with permission.)
FIGURE 16-19
annual propagation of new generations of cuttings-collected from asexually propagated container-produced woody and herbaceous ornamentals. 6. Establish "stoolshoot" beds by cutting shoots nearly to the ground (90). This is perhap one of the reasons for success in rooting in mound and trench layering (see Chapter 1Rejuvenation shift from the adult phase to the juvenile or transitional phase.